a Department of Biology , The University of York , York , UK.
Cell Cycle. 2018;17(18):2268-2283. doi: 10.1080/15384101.2018.1526600. Epub 2018 Oct 6.
CIZ1 promotes cyclin-dependent DNA replication and resides in sub-nuclear foci that are part of the protein nuclear matrix (NM), and in RNA assemblies that are enriched at the inactive X chromosome (Xi) in female cells. It is subjected to alternative splicing, with specific variants implicated in adult and pediatric cancers. CIZ1-F is characterized by a frame shift that results from splicing exons 8-12 leading to inclusion of a short alternative reading frame (ARF), excluding the previously characterized C-terminal NM anchor domain. Here, we apply a set of novel variant-selective molecular tools targeted to the ARF to profile the expression of CIZ1-F at both transcript and protein levels, with focus on its relationship with the RNA-dependent and -independent fractions of the NM. Unlike full-length CIZ1, CIZ1-F does not accumulate at Xi, though like full-length CIZ1 it does resist extraction with DNase. Notably, CIZ1-F is sensitive to RNase identifying it as part of the RNA-fraction of the NM. In quiescent cells CIZ1-F transcript expression is suppressed and CIZ1-F protein is excluded from the nucleus, with re-expression not observed until the second cell cycle after exit from quiescence. Importantly, CIZ1-F is over-expressed in common solid tumors including colon and breast, pronounced in early stage but not highly-proliferative late stage tumors. Moreover, expression was significantly higher in hormone receptor negative breast tumors than receptor positive tumors. Together these data show that CIZ1-F is expressed in proliferating cells in an unusual cell cycle-dependent manner, and suggest that it may have potential as a tumor biomarker.
CIZ1 促进细胞周期依赖性 DNA 复制,并定位于核基质(NM)亚核焦点和富含女性细胞中失活 X 染色体(Xi)的 RNA 组装体中。它经历可变剪接,特定变体与成人和儿科癌症有关。CIZ1-F 的特征是由于剪接外显子 8-12 导致包含短的替代阅读框(ARF)而导致的移码,排除了先前表征的 C 末端 NM 锚定结构域。在这里,我们应用一组针对 ARF 的新型变体选择性分子工具来分析 CIZ1-F 在转录本和蛋白质水平上的表达情况,重点是其与 NM 的 RNA 依赖性和非依赖性部分的关系。与全长 CIZ1 不同,CIZ1-F 不会在 Xi 处积累,尽管与全长 CIZ1 一样,它也不会抵抗 DNA 酶的提取。值得注意的是,CIZ1-F 对 RNase 敏感,表明它是 NM 的 RNA 部分的一部分。在静止细胞中,CIZ1-F 转录本表达受到抑制,CIZ1-F 蛋白从核内排出,直到静止后第二个细胞周期才重新表达。重要的是,CIZ1-F 在常见的实体瘤中过度表达,包括结肠癌和乳腺癌,在早期阶段明显,但在晚期高增殖性肿瘤中不明显。此外,在激素受体阴性的乳腺癌中表达显著高于受体阳性的肿瘤。这些数据表明,CIZ1-F 以一种不寻常的细胞周期依赖性方式在增殖细胞中表达,并表明它可能具有作为肿瘤生物标志物的潜力。